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'''Caucasus hunter-gatherer''' ('''CHG'''), also called '''Satsurblia cluster''',{{sfn|Fu|Posth|Hajdinjak|Petr|2016}}<ref>{{cite journal |author1=Eisenmann, S. |author2= Bánffy, E. |author3=van Dommelen, P. |display-authors=etal |year=2018 |title=Reconciling material cultures in archaeology with genetic data: The nomenclature of clusters emerging from archaeogenomic analysis |journal=Scientific Reports |volume=8 |issue= 1 |pages= 13003 |doi=10.1038/s41598-018-31123-z |pmid= 30158639 |pmc= 6115390 |bibcode= 2018NatSR...813003E }}</ref> is an [[anatomically modern human]] [[human genetics|genetic lineage]], first identified in a 2015 study,{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}{{sfn|Fu|Posth|Hajdinjak|Petr|2016}} based on the [[population genetics]] of several modern [[Eurasia|Western Eurasian]] ([[Europe]]an, [[Caucasus|Caucasian]] and [[Near Eastern]]) populations.<ref name="bbcnov16"/><ref name="Dutchen 2016">{{cite web|last1=Dutchen|first1=Stephanie|title=History on Ice|url=http://hms.harvard.edu/news/history-ice|publisher=[[Harvard Medical School]]|access-date=11 May 2016|date=2 May 2016}}</ref>
It represents an ancestry maximised in some [[Upper Paleolithic]] and [[Mesolithic]] hunter-gatherer groups in the [[Caucasus]]. These groups are also very closely related to Mesolithic hunter-gatherers and [[Neolithic]] farmers in the [[Iranian Plateau]], who are sometimes included within the CHG group.<ref>{{Cite journal |
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The CHG lineage is suggested to have diverged from the ancestor of [[Western Hunter-Gatherers]] (WHGs) probably during the [[Last Glacial Maximum]] (sometime between 45,000 to 26,000 years ago).<ref>{{cite journal |title='Fourth strand' of European ancestry originated with hunter-gatherers isolated by Ice Age |journal=University of Cambridge |date=16 November 2015 |url=https://www.cam.ac.uk/research/news/fourth-strand-of-european-ancestry-originated-with-hunter-gatherers-isolated-by-ice-age |language=en|quote=By reading the DNA, the researchers were able to show that the lineage of this fourth Caucasus hunter-gatherer strand diverged from the western hunter-gatherers just after the expansion of anatomically modern humans into Europe from Africa.}}</ref> They further separated from the [[Anatolian hunter-gatherers|Anatolian hunter-gatherer]] (AHG) lineage later, suggested to around 25,000 years ago during the late LGM period.{{sfn|Fu|Posth|Hajdinjak|Petr|2016}}<ref>{{Cite journal |last1=Marchi |first1=Nina |last2=Winkelbach |first2=Laura |last3=Schulz |first3=Ilektra |last4=Brami |first4=Maxime |last5=Hofmanová |first5=Zuzana |last6=Blöcher |first6=Jens |last7=Reyna-Blanco |first7=Carlos S. |last8=Diekmann |first8=Yoan |last9=Thiéry |first9=Alexandre |last10=Kapopoulou |first10=Adamandia |last11=Link |first11=Vivian |last12=Piuz |first12=Valérie |last13=Kreutzer |first13=Susanne |last14=Figarska |first14=Sylwia M. |last15=Ganiatsou |first15=Elissavet |date=May 2022 |title=The genomic origins of the world's first farmers |url=|journal=Cell |volume=185 |issue=11 |pages=1842–1859.e18 |doi=10.1016/j.cell.2022.04.008 |issn=0092-8674 |pmc=9166250 |pmid=35561686}}</ref> The Caucasus hunter-gatherers managed to survive in isolation since the late LGM period as a distinct population, and display high genetic affinities to Mesolithic and Neolithic populations on the Iranian plateau, such as Neolithic specimens found in [[Ganj Dareh]]. The CHG display higher genetic affinities to European and Anatolian groups than Iranian hunter-gatherers do, suggesting a possible cline and geneflow into the CHG and less into Mesolithic and Neolithic Iranian groups.{{sfn|Jones|Gonzalez-Fortes|Connell|Siska|2015}}<ref>{{Cite journal |last1=Gallego-Llorente |first1=M. |last2=Connell |first2=S. |last3=Jones |first3=E. R. |last4=Merrett |first4=D. C. |last5=Jeon |first5=Y. |last6=Eriksson |first6=A. |last7=Siska |first7=V. |last8=Gamba |first8=C. |last9=Meiklejohn |first9=C. |last10=Beyer |first10=R. |last11=Jeon |first11=S. |last12=Cho |first12=Y. S. |last13=Hofreiter |first13=M. |last14=Bhak |first14=J. |last15=Manica |first15=A. |date=9 August 2016 |title=The genetics of an early Neolithic pastoralist from the Zagros, Iran |journal=Scientific Reports |language=en |volume=6 |issue=1 |pages=31326 |doi=10.1038/srep31326 |pmid=27502179 |pmc=4977546 |bibcode=2016NatSR...631326G |issn=2045-2322}}</ref>
An alternative model without the need of significant amounts of ANE ancestry has been presented by Vallini et al. (2024), suggesting that the initial Iranian hunter-gatherer-like population which is basal to the CHG formed primarily from a deep Ancient West Eurasian lineage ('WEC2', {{Circa|72%}}), and from varying degrees of [[Ancient East Eurasians|Ancient East Eurasian]] ({{Circa|10%}}) and [[Basal Eurasian]] ({{Circa|18%}}) components. The Ancient West Eurasian component associated with Iranian hunter-gatherers (WEC2) is inferred to have diverged from the West Eurasian Core lineage (represented by [[Kostenki-14]]; WEC), with the WEC2 component staying in the region of the [[Iranian Plateau]], while the proper WEC component expanded into Europe.<ref>{{Cite journal |last1=Vallini |first1=Leonardo |last2=Zampieri |first2=Carlo |last3=Shoaee |first3=Mohamed Javad |last4=Bortolini |first4=Eugenio |last5=Marciani |first5=Giulia |last6=Aneli |first6=Serena |last7=Pievani |first7=Telmo |last8=Benazzi |first8=Stefano |last9=Barausse |first9=Alberto |last10=Mezzavilla |first10=Massimo |last11=Petraglia |first11=Michael D. |last12=Pagani |first12=Luca |date=25 March 2024 |title=The Persian plateau served as hub for Homo sapiens after the main out of Africa dispersal |journal=Nature Communications |language=en |volume=15 |issue=1 |pages=1882 |doi=10.1038/s41467-024-46161-7 |issn=2041-1723 |pmc=10963722 |pmid=38528002|bibcode=2024NatCo..15.1882V }}</ref>
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Some scholars argue that the archaic PIE ('[[Indo-Hittite|Indo-Anatolian']]) language may have originated among a CHG-rich population in [[West Asia|Western Asia]], based on the lack of EHG ancestry in the probable speakers of [[Anatolian languages]].<ref>{{Cite journal |last1=Lazaridis |first1=Iosif |last2=Alpaslan-Roodenberg |first2=Songül |last3=Acar |first3=Ayşe |last4=Açıkkol |first4=Ayşen |last5=Agelarakis |first5=Anagnostis |last6=Aghikyan |first6=Levon |last7=Akyüz |first7=Uğur |last8=Andreeva |first8=Desislava |last9=Andrijašević |first9=Gojko |last10=Antonović |first10=Dragana |last11=Armit |first11=Ian |last12=Atmaca |first12=Alper |last13=Avetisyan |first13=Pavel |last14=Aytek |first14=Ahmet İhsan |last15=Bacvarov |first15=Krum |date=26 August 2022 |title=The genetic history of the Southern Arc: A bridge between West Asia and Europe |journal=Science |language=en |volume=377 |issue=6609 |pages=eabm4247 |doi=10.1126/science.abm4247 |issn=0036-8075 |pmc=10064553 |pmid=36007055}}</ref> Others, such as Anthony, suggest that PIE was spoken by EHGs living in [[Eastern Europe]].<ref>{{Cite journal |last=Anthony |first=David |date=1 January 2019 |title=Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard |url=https://www.academia.edu/39985565 |journal=Journal of Indo-European Studies}}</ref>
According to Jones ''et al''. (2015), Caucasus hunter-gatherer (CHG) "genomes significantly contributed to the Yamnaya steppe herders who migrated into Europe ~3,000 BCE, supporting a formative Caucasus influence on this important Early [[Bronze Age]] culture. CHG left their imprint on modern populations from the [[Caucasus]] and also [[Central Asia|Central]] and [[South Asia]] possibly correlating with the arrival of [[Indo-Aryan language]]s."<ref>{{harvnb|Jones|Gonzalez-Fortes|Connell|Siska|2015}}: "Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers ~45 kya, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers ~25 kya, around the Last Glacial Maximum."</ref> For example, about 50–70% of Armenian ancestry is derived from CHG, persisting from Neolithic times to the present.<ref>{{Cite journal |last1=Dergachev |first1=Valentin |last2=Shephard |first2=Henry |last3=Sirbu |first3=Ghenadie |last4=Szécsényi-Nagy |first4=Anna |date=2022 |title=The genetic history of the Southern Arc: A bridge between West Asia and Europe |url=https://www.researchgate.net/publication/363157308 |journal=Science |volume=377 |issue=6609 |pages=982–987 |via=ResearchGate}}</ref> Wang et al. (2018) analysed genetic data of the North Caucasus of fossils dated between the 4th and 1st millennia BC and found correlation with modern groups of the South Caucasus, concluding that "unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement".{{sfn|Wang|Reinhold|Kalmykov|Wissgott|2018}} According to Allentoft ''et al.'' (2024), The arrival and admixture of CHG with Caspian steppe cultures is dated to about 7,300-years-old, which is seen in two ancient samples from Golubaya Krinitsa with 18–24% admixture.<ref name="Allentoft_2024">{{Cite journal |last1=Allentoft |first1=Morten E. |last2=Sikora |first2=Martin |last3=Refoyo-Martínez |first3=Alba |last4=Irving-Pease |first4=Evan K. |last5=Fischer |first5=Anders |last6=Barrie |first6=William |last7=Ingason |first7=Andrés |last8=Stenderup |first8=Jesper |last9=Sjögren |first9=Karl-Göran |last10=Pearson |first10=Alice |last11=Sousa da Mota |first11=Bárbara |last12=Schulz Paulsson |first12=Bettina |last13=Halgren |first13=Alma |last14=Macleod |first14=Ruairidh |last15=Jørkov |first15=Marie Louise Schjellerup |date=January 2024 |title=Population genomics of post-glacial western Eurasia
== Ancient Greece, Aegean and Italy ==
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Genetic analysis shows that Iranian-related ancestry, which was widespread in the Aegean by the Middle Bronze Age in association with the Minoan and Mycenaean cultures, had also spread as far west as Sicily in substantial proportion at least by the time of the Mycenaeans. One possibility is this ancestry spread west along with the Mycenaean cultural expansion.<ref>{{Cite journal |last1=Fernandes |first1=Daniel M. |last2=Mittnik |first2=Alissa |last3=Olalde |first3=Iñigo |last4=Lazaridis |first4=Iosif |last5=Cheronet |first5=Olivia |last6=Rohland |first6=Nadin |last7=Mallick |first7=Swapan |last8=Bernardos |first8=Rebecca |last9=Broomandkhoshbacht |first9=Nasreen |last10=Carlsson |first10=Jens |last11=Culleton |first11=Brendan J. |last12=Ferry |first12=Matthew |last13=Gamarra |first13=Beatriz |last14=Lari |first14=Martina |last15=Mah |first15=Matthew |date=March 2020 |title=The Spread of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean |journal=Nature Ecology & Evolution |volume=4 |issue=3 |pages=334–345 |doi=10.1038/s41559-020-1102-0 |issn=2397-334X |pmc=7080320 |pmid=32094539|bibcode=2020NatEE...4..334F }}</ref> An arrival of the CHG-related component in Southern Italy from the Southern part of the Balkan Peninsula, including the Peloponnese, is compatible with the identification of genetic corridors linking the two regions and the presence of Southern European ancient signatures in Italy.<ref>{{Cite journal |last1=Raveane |first1=A. |last2=Aneli |first2=S. |last3=Montinaro |first3=F. |last4=Athanasiadis |first4=G. |last5=Barlera |first5=S. |last6=Birolo |first6=G. |last7=Boncoraglio |first7=G. |last8=Di Blasio |first8=A. M. |last9=Di Gaetano |first9=C. |last10=Pagani |first10=L. |last11=Parolo |first11=S. |last12=Paschou |first12=P. |last13=Piazza |first13=A. |last14=Stamatoyannopoulos |first14=G. |last15=Angius |first15=A. |date=4 September 2019 |title=Population structure of modern-day Italians reveals patterns of ancient and archaic ancestries in Southern Europe |journal=Science Advances |volume=5 |issue=9 |pages=eaaw3492 |doi=10.1126/sciadv.aaw3492 |issn=2375-2548 |pmc=6726452 |pmid=31517044|bibcode=2019SciA....5.3492R }}</ref> Collected data from Iron Age individuals dating from 900 to 200 BCE (including the Republican period) group shows a clear ancestry shift from the Copper Age, interpreted by ADMIXTURE as the addition of a Steppe-related ancestry component, and an increase in the Neolithic-Iranian component.<ref>{{Cite journal |last1=Antonio |first1=Margaret L. |last2=Gao |first2=Ziyue |last3=Moots |first3=Hannah M. |last4=Lucci |first4=Michaela |last5=Candilio |first5=Francesca |last6=Sawyer |first6=Susanna |last7=Oberreiter |first7=Victoria |last8=Calderon |first8=Diego |last9=Devitofranceschi |first9=Katharina |last10=Aikens |first10=Rachael C. |last11=Aneli |first11=Serena |last12=Bartoli |first12=Fulvio |last13=Bedini |first13=Alessandro |last14=Cheronet |first14=Olivia |last15=Cotter |first15=Daniel J. |date=8 November 2019 |title=Ancient Rome: A genetic crossroads of Europe and the Mediterranean |journal=Science |volume=366 |issue=6466 |pages=708–714 |doi=10.1126/science.aay6826 |issn=0036-8075 |pmc=7093155 |pmid=31699931|bibcode=2019Sci...366..708A }}</ref>
== Central Asia and
CHG/Iranian Plateau Neolithic-like ancestry is prominent in pre-steppe admixture [[Chalcolithic]] and [[Bronze Age]] (4500–2000 BCE) populations in Central Asia, like the [[Bactria–Margiana Archaeological Complex]] (which also had [[Anatolian Neolithic Farmers|Anatolian Neolithic Farmer]]-related ancestry)<ref name=":3" /> as well as in the northwestern Indian subcontinent such as in sites in or adjacent to the [[Indus Valley Civilisation]] (who have mixed CHG-related and [[Ancient Ancestral South Indian]] ancestry).<ref name=":2" /><ref name=":3" /> It is unclear as to whether the dispersal of CHG/Iranian Plateau-related ancestry eastwards to the Indian subcontinent was the result of the migration of farmers or an earlier dispersal of hunter-gatherers who later adopted farming, but this dispersal likely occurred sometime before 6000 BCE due to the lack of Anatolian Farmer-related ancestry in ancient South Asians, but which is present in the Iranian Plateau after this time. This pre-steppe CHG-related ancestry makes up a significiant proportion of the ancestry of modern South Asians.<ref name=":3" /> WSHs, who were of significant CHG ancestry, also later migrated into Central Asia and
==See also==
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